Pollen morphology, deep learning, phylogenetics, and the evolution of environmental adaptations in Podocarpus.

Podocarpus pollen morphology is shaped by both phylogenetic history and the environment. We analyzed the relationship between pollen traits quantified using deep learning and environmental factors within a comparative phylogenetic framework. We investigated the influence of mean annual temperature, annual precipitation, altitude, and solar radiation in driving morphological change. We used trait-environment regression models to infer the temperature tolerances of 31 Neotropical Podocarpidites fossils. Ancestral state reconstructions were applied to the Podocarpus phylogeny with and without the inclusion of fossils. Our results show that temperature and solar radiation influence pollen morphology, with thermal stress driving an increase in pollen size and higher ultraviolet B radiation selecting for thicker corpus walls. Fossil temperature tolerances inferred from trait-environment models aligned with paleotemperature estimates from global paleoclimate models. Incorporating fossils into ancestral state reconstructions revealed that early ancestral Podocarpus lineages were likely adapted to warm climates, with cool-temperature tolerance evolving independently in high-latitude and high-altitude species. Our results highlight the importance of deep learning-derived features in advancing our understanding of plant environmental adaptations over evolutionary timescales. Deep learning allows us to quantify subtle interspecific differences in pollen morphology and link these traits to environmental preferences through statistical and phylogenetic analyses.